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Light Microscopic and Scanning Electron Microscopic Techniques to Characterize Nutlets of Some Indian Cyperoideae (Cyperaceae) and Their Taxonomic Significance
Abstract
In the present investigation, nutlet morphological and micro‐morphological characters were analyzed using Light Microscope (LM) and Scanning Electron Microscope (SEM) in 38 taxa under 13 genera from the sub‐family Cyperoideae of Cyperaceae to find out whether these characters are taxonomically important or not. Nutlet morphology and surface characters of the representative taxa from all the possible tribes under the sub‐family Cyperoideae namely, Fuireneae, Cypereae, Cariceae, Abildgaardieae, Eleocharideae, Pseudoschoeneae, Schoenoplecteae, and Sclerieae were evaluated for their taxonomic significance. Cluster analysis was employed considering nutlet morphological characters to determine the overall similarity among the taxa based on 153 character states. Except in few specified cases, grouping of the taxa in the clusters is in accordance to the taxonomic treatments made by recent Cyperologists. Nutlets in the tribe Abildgaardieae showed maximum level of variability in size, shape, and surface ornamentation at the level of higher taxon, but showed specificity at the species level. Tuberculate, striate‐reticulate, and transversely wavy ridged surface ornamentations were found in different species of Fimbristylis . Previously described cryptic variation, and effect of different ploidy level were not reflected in nutlet surface morphology and micromorphology in Fimbristylis dichotoma and F. ovata complexes which was previously made the taxa taxonomically very difficult. Serrulate anticlinal wall in F. bisumbellata was the most unique in Fimbristylis . Species under the megadiverse genus Carex representing the tribe Cariceae showed very unique type of surface ornamentation. Surface walls of all the studied species of Carex were characterized by polygonal epidermal cells with single conical silica body (2–3 per cell in C. speciosa ) of variable length and sizes. Most interestingly, in C. nubigena , presence of the central silica body and peripheral satellites was not consistent. Based on the presence and absence, two different variants under the species were identified. In C. nubigena , when silica body was present, epidermal cells were characterized by central conical silica body surrounded by variable number of satellites. The present investigation first time reports this novel nutlet surface character in C. nubigena . Among the studied characters, length and height of conical, height of apex, and width of apex are variable among Carex species. On the other hand, exclusion of the genera Schoenoplectiella and Schoenoplectus from tribe Scirpeae s.l. and placement under the tribes Pseudoschoeneae and Schoenoplecteae (respectively) was also supported by the present investigation. The present study also confirms that nutlet morphological and micro‐morphological characters are useful in identification and arrangement of different taxa under the subfamily Cyperoideae of Cyperaceae. Result of the present investigation was correlated and discussed in comparative manner with the treatments of the recent past.
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Carex linanensis X.D.Qiu & X.F.Jin, a new species in sect. Mitratae of the sedge family (Cyperaceae) from north-western Zhejiang is described and illustrated. We performed a statistical comparison of the new species with other closely-related species from the same section. Carex linanensis is similar to Carex sachalinensis F.Schmidt, but differs in having leaf blades 1–2 mm wide (vs. 2.5–3.5 mm wide), utricles longer than pistillate glumes, with beak margin smooth (vs. barbate) and peduncles of lateral spikes enclosed in bract sheaths (vs. exserted from bract sheaths).
This paper provides a linear classification of two subfamilies, 24 tribes, 10 subtribes and 95 genera of the monocot family Cyperaceae (Poales), based on a stable phylogenetic framework resulting from years of morphological, molecular phylogenetic and phylogenomic studies. The family includes c. 5687 species. The most species-rich tribes are the monogeneric tribe Cariceae with c. 2003 species, and tribe Cypereae with c. 1131 species. The highest generic diversity is found in tribe Schoeneae (25 genera), which resulted in the recognition of eight subtribes to facilitate studying this group. The linear classification will help the organisation of Cyperaceae specimens in herbaria according to a systematic order and provides an easy-to-use summary of the current classification of the family.
Carex sect. Confertiflorae s.l. is a medium sized species group (ca. 40 species) with its center of diversity in E Asia (China and Japan). According to morphological traits, the section has been proposed to split into two sections (sects. Confertiflorae sensu Ohwi and Molliculae Ohwi) up to five different ones (sects. Confertiflorae s.s., Molliculae, Dispalatae Ohwi, Ischnostachyae Ohwi and Alliiformes Akiyama). Recent phylogenetic reconstructions showed Confertiflorae s.l. not to be monophyletic, since species traditionally considered part of it were found to belong to other clades, while species traditionally ascribed to other sections were nested within it. In this study, we investigated the phylogenetic structure, morphological affinities, and biogeographic history of sect. Confertiflorae s.l. We performed a taxon-based approach to explore the morphological affinities of the species considered in sect. Confertiflorae and compared the micromorphology of the nutlets of almost all the taxa using SEM. We included 40 samples representing 31 species/subspecies of sect. Confertiflorae s.l., and used two nuclear (ETS and ITS) and three plastid (trnL-F, matK and rpl32-trnLUAG) molecular markers to reconstruct the phylogeny of the group. The phylogenetic analyses confirmed the polyphyly of sect. Confertiflorae s.l., whose representatives were found within five distinct clades. From these, two clades, sect. Confertiflorae and sect. Molliculae were found to be closely related and contained the majority of the species. The composition of the two clades agreed with the morphological structure of the group, and we confirmed an exclusive combination of features (viz. color of basal sheaths, presence of bract sheath, peduncle of lowest spike, inflorescence sex distribution, shape of pistillate glume apex, and color and veins of utricle, among others) characterizing each of the two clades. The origin of the two clades was found to be in the early Pliocene while the majority of the diversification events within each clade took place during the Pleistocene. This illustrate that despite Asia has been regarded as having little potential ecological space for Carex to diversify because its climate stability, groups of sedges sub-endemic from that area may have a fairly recent origin related to glaciations. We proposed the rearrangement of sect. Confertiflorae as previously conceived as three independent sections: the monotypic Alliiformes, sect. Molliculae, and sect. Paludosae.
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Cyperaceae (sedges) are the third largest monocot family and are of considerable economic and ecological importance. Sedges represent an ideal model family to study evolutionary biology because of their species richness, global distribution, large discrepancies in lineage diversity, broad range of ecological preferences, and adaptations including multiple origins of C4 photosynthesis and holocentric chromosomes. Goetghebeur’s seminal work on Cyperaceae published in 1998 provided the most recent complete classification at tribal and generic level, based on a morphological study of Cyperaceae inflorescence, spikelet, flower and embryo characters plus anatomical and other information. Since then, several family‐level molecular phylogenetic studies using Sanger sequence data have been published. Here, more than 20 years after the last comprehensive classification of the family, we present the first family‐wide phylogenomic study of Cyperaceae based on targeted sequencing using the Angiosperms353 probe kit sampling 311 accessions. Additionally, 62 accessions available from GenBank were mined for overlapping reads and included in the phylogenomic analyses. Informed by this backbone phylogeny, a new classification for the family at the tribal, subtribal and generic levels is proposed. The majority of previously recognized suprageneric groups are supported, and for the first time we establish support for tribe Cryptangieae as a clade including the genus Koyamaea. We provide a taxonomic treatment including identification keys and diagnoses for the 2 subfamilies, 24 tribes and 10 subtribes and basic information on the 95 genera. The classification includes five new subtribes in tribe Schoeneae: Anthelepidinae, Caustiinae, Gymnoschoeninae, Lepidospermatinae and Oreobolinae.
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Cyperaceae in the campos de natureza of Cametá, Pará, Amazon, Brazil. This study presents a taxonomic treatment of genera and
species of Cyperaceae Juss. in the campos de natureza of Cameta, Pará, Amazon, Brazil. After intense field expeditions and studies of herbarium collections, 11 genera and 51 species of Cyperaceae were identified for the area: Rhynchospora Vahl (17 species), Scleria P. J. Bergius (9), Cyperus L. (7), Eleocharis R. Br. (4), Bulbostylis Kunth (3), Lagenocarpus Nees (3), Fimbristylis Vahl (3), Calyptrocarya Nees (2), Diplacrum R. Br. (1), Fuirena Rottb. (1) and Hypolytrum Rich (1). Four species are recorded in the state of Pará for the first time: Eleocharis angustispicula R. Trevis., Eleocharis jelskiana Boeck., Rhynchospora curvula Griseb., and Scleria amazonica Camelb., M. T. Strong & Goetgh. A taxonomic identification key, morphological descriptions, taxonomic comments, geographic distribution, habitat data and illustrations for all species nutlets are provided.
Molecular phylogenetic studies based on Sanger sequences have shown that Cyperaceae tribe Fuireneae s.l. is paraphyletic. However, taxonomic sampling in these studies has been poor, topologies have been inconsistent, and support for the backbone of trees has been weak. Moreover, uncertainty still surrounds the morphological limits of Schoenoplectiella, a genus of mainly small, amphicarpic annuals that was recently segregated from Schoenoplectus. Consequently, despite ample evidence from molecular analyses that Fuireneae s.l. might consist of two to four tribal lineages, no taxonomic changes have yet been made. Here, we use the Angiosperms353 enrichment panel for targeted sequencing in order to: (1) clarify the relationships of Fuireneae s.l. with the related tribes Abildgaardieae, Eleocharideae and Cypereae; (2) define the limits of Fuireneae s.s., and (3) test the monophyly of Fuireneae s.l. genera with emphasis on Schoenoplectus and Schoenoplectiella. Using more than a third of Fuireneae s.l. diversity, our phylogenomic analyses strongly support six genera and four major Fuireneae s.l. clades that we recognise as tribes: Bolboschoeneae stat.nov., Fuireneae s.s., Schoenoplecteae, and Pseudoschoeneae tr.nov. These results are consistent with morphological, micromorphological (nutlet epidermal cell shape), and embryo differences detected for each tribe. At the generic level, most sub‐Saharan African perennials currently treated in Schoenoplectus are transferred to Schoenoplectiella. Our targeted sequencing results show that these species are nested in Schoenoplectiella, and their treatment here is consistent with micromorphological and embryo characters shared by all Schoenoplectiella species. Keys to recognised tribes and genera are provided.
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Citation: Rewicz A, Marciniuk J, Marciniuk P (2020) Achene micromorphology and its taxonomic significance in some species in Taraxacum sect. Palustria (Asteraceae). PhytoKeys 166: 1-28. https://doi. Abstract The genus Taraxacum is one of the largest and taxonomically most complicated apomictic genera. Currently , it is considered to consist of over 2800 species placed within 60 sections. Due to the large number of species, and their uniform morphological design and plasticity of leaves, the identification of plants at the species level is challenging even for specialists. This problem significantly hinders the study of their properties and the rational use of these valuable medicinal and nutritional plants. This paper presents the results of research on the morphology and micromorphology of achenes of 28 Taraxacum species of sect. Palustria and for comparison one species per section of: Erythrosperma, Naevosa, Piesis, and Taraxacum. The achenes were measured with a stereoscopic microscope and a biometric program, and micromorpho-logical studies were performed by scanning electron microscopy. It has been shown that traits associated with achene morphology and micromorphology have a high diagnostic value, allowing us to distinguish sections as well as species within the sect. Palustria. Based on the examined achene features, a dichotomous key for determining the studied species was constructed.
Nutlets of 13 taxa of subfamily Cyperoideae (Cyperaceae) from Egypt were investigated using light and scanning electron
microscopes to ascertain the taxonomic utility of the nutlet in corroborating and elaborating on distinctions between closely related
genera and species. The studied taxa included representatives of tribes Fuireneae, Cypereae, Cariceae, Abildgaardieae and Schoeneae.
Nutlet shape, size, color, surface, epidermal cell shape, anticlinal boundaries, outer periclinal cell wall and relief of outer cell walls were
provided. It was found that the epidermis has useful microscopic characters to help in the taxonomic assessment of Cyperoideae. Seven
different types of anticlinal cell wall boundaries were described, four types of outer periclinal cell walls, and five different forms of the
secondary cell walls were recognized. A diagnostic key for the studied taxa based on nutlet characters was presented. The results showed
congruence between the UPGMA dendrogram and principal component analysis in suggesting two major groups and six subgroups.
The most important characters with the highest factor loadings along the first two PCA axes included nutlet shape, nutlet apex, surface
topography and the presence of silica bodies. The results of this study confirmed the heterogeneity of the Cyperoideae and indicated that
Fuireneae is the most heterogeneous tribe.
This study aims to contribute to the knowledge on the taxonomy of Scleria in sandy coastal plains (restinga) in Pará state, Brazil. Material deposited in the herbaria of Museu Paraense Emílio Goeldi (MG), Embrapa Amazônia Oriental (IAN), and Instituto de Estudos Costeiros da Universidade Federal do Pará (HBRA) were analyzed and species were identified using relevant literature, protologues, and types available online. We recorded nine species of Scleria for restinga in the state: Scleria bracteata, S. gaertneri, S. hirtella, S. interrupta, S. macrophylla, S. microcarpa, S. reticularis, S. secans, and S. violacea. An identification key, descriptions, illustrations, and comments about diagnostic characters and geographic distribution are provided for these species.
The megadiverse genus Carex (c. 2000 species, Cyperaceae) has a nearly cosmopolitan distribution, displaying
an inverted latitudinal richness gradient with higher species diversity in cold‐temperate areas of the Northern
Hemisphere. Despite great expansion in our knowledge of the phylogenetic history of the genus and many molecular
studies focusing on the biogeography of particular groups during the last few decades, a global analysis of Carex
biogeography and diversification is still lacking. For this purpose, we built the hitherto most comprehensive Carex‐dated
phylogeny based on three markers (ETS–ITS–matK), using a previous phylogenomic Hyb‐Seq framework, and a sampling
of two‐thirds of its species and all recognized sections. Ancestral area reconstruction, biogeographic stochastic mapping,
and diversification rate analyses were conducted to elucidate macroevolutionary biogeographic and diversification
patterns. Our results reveal that Carex originated in the late Eocene in E Asia, where it probably remained until the
synchronous diversification of its main subgeneric lineages during the late Oligocene. E Asia is supported as the cradle of
Carex diversification, as well as a “museum” of extant species diversity. Subsequent “out‐of‐Asia” colonization patterns
feature multiple asymmetric dispersals clustered toward present times among the Northern Hemisphere regions, with
major regions acting both as source and sink (especially Asia and North America), as well as several independent
colonization events of the Southern Hemisphere. We detected 13 notable diversification rate shifts during the last 10My, including remarkable radiations in North America and New Zealand, which occurred concurrently with the late Neogene
global cooling, which suggests that diversification involved the colonization of new areas and expansion into novel areas
of niche space.
A taxonomic revision of the genus Scleria (Cyperoideae, Cyperaceae) in Madagascar is presented. Herbarium specimens have been examined and 422 identified to species level. Our results recognise 25 species of Scleria from Madagascar, plus an additional heterotypic variety. Eight species are endemic to Madagascar, two are near endemic, eight taxa are also found on mainland Africa, and eight are widespread tropical taxa. Scleria achtenii is reported from Madagascar for the first time, and S. rosea is accepted at species level instead of being considered as a synonym of S. trialata. Distribution maps, conservation assessments, and notes on synonymy, ecology and ethnobotany are provided. Fourty-seven names are typified. Three rare endemic species: S. andringitrensis, S. madagascariensis and S. perpusilla, are assessed as threatened; and a recently described species, S. ankaratrensis, is indicated as Data Deficient. The most species-rich infrageneric taxa, sections Hypoporum, Abortivae and Foveolidia include 18 taxa in total, and showed strong differences in habitat preference.
Cyperaceae tribe Fuireneae consists of six genera (Actinoscirpus, Pseudoschoenus, Fuirena, Bolboschoenus, Schoenoplectus, and Schoenoplectiella) and approximately 155 species distributed widely across all continents except Antarctica. Recent molecular analyses suggest that Fuireneae is paraphyletic with respect to Cypereae, but limited taxonomic sampling and low branch support have resulted in unresolved relationships among key genera. To address the potential paraphyly of Fuireneae as well as intratribal and intertribal relationships, thirty taxa representing major lineages from the six Fuireneae genera within Cypereae were sequenced for five coding and non-coding regions from the nuclear (internal transcribed spacer, ITS) and chloroplast (trnL intron/ trnL-trnF spacer; partial ndhF, rbcL and matK genes) genomes. Maximum parsimony, maximum likelihood and Bayesian analyses of the dataset strongly support Fuireneae as a grade of three clades: (1) Fuirena, (2) Bolboschoenus and (3) a group composed of Cypereae and the four remaining Fuireneae genera (Schoenoplectus, Schoenoplectiella, Actinoscirpus, and Pseudoschoenus). Individual and combined gene trees consistently position Isolepis humillima within a well supported clade consisting of species in Schoenoplectiella sect. Schoenoplectiella. Based on molecular evidence, we transfer Isolepis humillima to Schoenoplectiella humillima, and suggest that a large-scale revision of tribal limits is required to reflect the natural relationships of the genera currently treated in Fuireneae.
Achenes from 23 specimens of Fimbristylis were examined using SEM to determine the distinguishing micromorphological characteristics that would be useful in establishing taxonomic relationships among the species populations. Various epidermal features were found to be useful for classifying specimens and clarifying the phylogenetic relationship between species and support species recognition. Achene micromorphology provides significant evidence that is useful in establishing new species within the genus and in revaluating of section or series realignment. Polyploid taxa could not be separated on the basis of achene micromorphology.
Achenes of roses were rarely studied and the studies were focused on anatomical research, mainly on pericarp structure and its development. We investigated the achene morphology by light- and scanning electron microscopy (LM and SEM) of 17 Rosa taxa from three sections (R. gallica from Gallicanae section, R. pendulina and R. rugosa from Rosa section, and R. agrestis, R. canina, R. canina var. corymbifera, R. dumalis, R. dumalis var. caesia, R. inodora, R. jundzillii, R. micrantha, R. rubiginosa, R. sherardii, R. ×subcanina, R. tomentosa, R. villosa, R. zalana from Caninae section). Eight quantitative and eight qualitative features were examined based on 9181 achenes, in total. Average achene size ranged from 4.37 to 5.39 mm in length and from 2.57 to 3.32 mm in width. The lowest morphological variability among the examined taxa was found in R. canina var. corymbifera, and the highest in R. gallica, R. inodora, and R. sherardii. The most diagnostic features of the achenes studied were suture (visible or invisible), presence or absence of hairs, hairs distribution and density, the exocarp sculpture and cuticle pattern type (we have distinguished four exocarp sculpture and three cuticle pattern types), and length. Qualitative achene features have significantly higher diagnostic value than quantitative ones. Taxonomical value of these features is quite high on the species and section level. Our study has shown that the previously mentioned morphological features of achenes can be used as valuable, additional diagnostic features in delimitation of Rosa taxa at the species and section level. Based on the morphological features of achenes, a determination key for all Rosa taxa studied was created.
The seed morphology of six species of the genus Juncus L. (Juncaceae), involving wild annuals and perennials was studied in Northeast of Iran using Light and Scanning Electron Microscope (SEM). Seed morphology (light microscope) of the investigated species exhibits certain variations in shape, size and surface ornamentation. Results of SEM analysis reveals that there are different patterns of seed surface. Seed shape and surface structure exhibit some diversity among taxa indicating systematic significance in species differentiation. An identification key to the investigated taxa is provided based on seed characters in this research.
Carex (sect. Rhomboidales) jubozanensis J. Oda & A. Tanaka is described as a new species from the western and central Honshu, Japan. This new species is characterized and clearly distinguished from its allies in the sect. Rhomboidales by the actively stoloniferous habit and the emarginate scales of both staminate and pistillate flowers. The micromorphology of achene surface is also different from that of its allies.
The structures of achene epidermal cells and their silica contents in ten species of Carex are illustrated and described here for the first time using scanning electron microscopy (SEM). The species examined in this study exhibited a wide variety of achene epidermal features, and the epidermal patterns of each taxon were distinctive. Achene micromorphology in the genus Carex can be used as a taxonomic criterion, but it should be combined with other features when diagnostic determinations of species are carried out.
A taxonomic study of Eleocharis acutangula (Roxb.) Schult, was conducted in order to better define this poorly understood and variable pantropical species. Multivariate statistical analysis, and ecological and distributional data of worldwide collections of E. acutangula provided the basis for its segregation into E. acutangula subsp. acutangula, E. acutangula subsp. breviseta D.J. Rosen, subsp. nov., and E. acutangula subsp. neotropica D.J. Rosen, subsp. nov. Nomenclatural research necessitated the lectotypification of E. acutangula and a heterotypic synonym, E. fistulosa Schult, var. robusta Boeck. A taxonomic treatment of E. acutangula is provided that includes a key to the subspecies, detailed descriptions, illustrations, and notes on habitat and distribution.
A new species of Carex sect. Rhomboidales, C. jianfengensis, is described and illustrated from Hainan, China. The new species is similar to C. zunyiensis but differs in having involucral bracts sparsely hispid and with ca.1 cm long sheaths; inflorescence with 4 spikes, terminal spike ca. 2.5 cm long, lateral spikes 2-3.5 × 0.7-1 cm; staminate glumes narrowly ovate, ca. 5 mm; pistillate glumes triangular-lanceolate, 5-7 mm; perigynia 6-8 × 3 mm and pubescent on veins; nutlet 4-5 mm long, rhombic-ovoid, trigonous, base with shortly stipitate, apex abruptly contracted into a erect short beak, and not expanding into an annulate orifice.
Abstract— Relationships within Schoenoplectus and Schoenoplectiella are largely unknown and the phylogenetic positions of these genera relative to the other four genera in Fuireneae and clade of Cypereae are unclear. A few studies with sparse or localized sampling have added valuable insights, but a North American sampling and a broad global perspective are needed. To generate a more robust phylogenetic hypothesis, we increased the number and breadth of taxon sampling in Schoenoplectus and Schoenoplectiella, including all constituent species in North America, all genera in Fuireneae, and strategically sampled genera in Cypereae. Phylogenetic relationships were estimated using DNA sequences from the nuclear ribosomal ITS region, chloroplast DNA trnL intron and trnL-trnF intergenic spacer region, and partial chloroplast DNA ndhF coding region and parsimony, likelihood, and Bayesian analyses. The proposed phylogeny reveals Pseudoschoenus, Schoenoplectiella, and Cypereae are supported as a clade, and Schoenoplectiella is paraphyletic and sister to Pseudoschoenus. Schoenoplectus is monophyletic, sister to Actinoscirpus. Schoenoplectus sections Schoenoplectus and Malacogeton were resolved; comprehensive sampling in Schoenoplectus section Schoenoplectus and unclear placement of S. californicus suggests the need to examine formerly recognized section Pterolepis. The proposed phylogeny supports the erection of sections in Schoenoplectiella, but indicates further morphological and molecular data is needed for section diagnoses. Two Cypereae taxa previously resolved in a Schoenoplectiella clade were included in this analysis: Scirpoides varia resolved in a clade with Cypereae taxa, and Isolepis humillima resolved within Schoenoplectiella. Results from the phylogenetic hypotheses support a need to revisit the generic placement of Isolepis humillima and revise Fuireneae to resolve tribal paraphyly.
Recent molecular phylogenetic analyses showed that Lipocarpha and Volkiella are nested in a paraphyletic Cyperus s.s. and therefore should be viewed as part of a broadly circumscribed genus Cyperus (Cyperaceae). In this paper, molecular phylogenetic analyses of Lipocarpha and Volkiella based on nuclear ribosomal ETS1f and plastid rpl32-trnL and trnH- psbA markers are presented. Separate gene trees as well as a species tree were constructed. Results indicate a polyphyletic Lipocarpha s.l. consisting of a paraphyletic core Lipocarpha s.s in which the monotypic Volkiella is included, and a small non-related clade with species formerly placed in the genus Rikliella. Core Lipocarpha s.s. encompasses six clades, which can be distinguished based on morphological characters. Floral developmental data for Lipocarpha rehmannii (the type of Rikliella) confirms that this species is not a true Lipocarpha s.s. Based on our findings, Lipocarpha s.l. and Volkiella are here included in Cyperus subg. Cyperus. New names and combinations for Lipocarpha s.l. and Volkiella species and a new sectional classification for these species are proposed.
A comprehensive, but simple-to-use software package for executing a range of standard numerical analysis and operations used in quantitative paleontology has been developed. The program, called PAST (PAleontological STatistics), runs on standard Windows computers and is available free of charge. PAST integrates spreadsheettype data entry with univariate and multivariate statistics, curve fitting, time-series analysis, data plotting, and simple phylogenetic analysis. Many of the functions are specific to paleontology and ecology, and these functions are not found in standard, more extensive, statistical packages. PAST also includes fourteen case studies (data files and exercises) illustrating use of the program for paleontological problems, making it a complete educational package for courses in quantitative methods.
Morphological description of seeds is a required step for analysis of biodiversity in natural populations and may
give clues to adaptive strategies in species evolution. The cardioid is the curve described by a point of one circumference
rolling around another circumference of equal radius. Models based on adjustment of seed shape
with cardioid curves have been described for Arabidopsis thaliana and the model legumes Lotus japonicus and
Medicago truncatula. In this work the model is applied to analyze seed morphology in populations of two subspecies
of Capparis spinosa growing in Tunisia. Adjustment of seed images to cardioid curves, followed by statistical
analysis of similarity in the complete images as well as in each of four quadrants, allows an accurate
description of seed shape. The results show differences in morphology between subspecies. Seeds of subsp.
rupestris present higher diversity of shape than seeds of subsp. spinosa. This may indicate primitiveness of C.
subsp. rupestris seeds, associated with nonspecialization. The results are discussed in relation to the ecological
strategies of both subspecies in their evolution.
Premise of the study:
Despite growing interest in the systematics and evolution of the hyperdiverse genus Carex, few studies have focused on its evolution using an absolute time framework. This is partly due to the limited knowledge of the fossil record. However, Carex fruits are not rare in certain sediments. We analyzed carpological features of modern materials from Carex sect. Phacocystis to characterize the fossil record taxonomically.
Methods:
We studied 374 achenes from modern materials (18 extant species), as well as representatives from related groups, to establish the main traits within and among species. We also studied 99 achenes from sediments of living populations to assess their modification process after decay. Additionally, we characterized 145 fossil achenes from 10 different locations (from 4-0.02 mya), whose taxonomic assignment we discuss.
Key results:
Five main characters were identified for establishing morphological groups of species (epidermis morphology, achene-utricle attachment, achene base, style robustness, and pericarp section). Eleven additional characters allowed the discrimination at species level of most of the taxa. Fossil samples were assigned to two extant species and one unknown, possibly extinct species.
Conclusions:
The analysis of fruit characters allows the distinction of groups, even up to species level. Carpology is revealed as an accurate tool in Carex paleotaxonomy, which could allow the characterization of Carex fossil fruits and assign them to subgeneric or sectional categories, or to certain species. Our conclusions could be crucial for including a temporal framework in the study of the evolution of Carex.
Micromorphology of the achene surface of 26 Brazilian species of Eleocharis was studied by scanning electron microscopy in order to evaluate its usefulness in the taxonomy of the genus. The results point out two pat-terns of cell organization according to silica structures of achenes. The first corresponds to a group of species (group A) that have small to medium cells arranged vertically. The second is found in those species (group B) with medium to large cells arranged horizontally. These data were useful in separating species of Eleocharis subgenus Scirpidium and E. subgenus Limnochloa (group B) from E. subgenus Eleocharis (group A). However, group A shows consider-able variation in silica wall arrangement. Eleocharis squamigera, previously considered as part of E. subgenus Eleocharis, shows features rather similar to those of Sci-rpidium, confirming recent phylogenies. The subgenus Limnochloa was clearly distinguished from others by achenes with large cells (over 55 lm width), presence of crenate or repand anticlinal walls, and some orifices near the wall in some species. The silica wall ornamentation seems to be a useful morphological tool for studying relationships between subgenera and distinguishes Limno-chloa from the other subgenera.
Descriptions of the 122 genera of Cyperaceae were automatically converted from a DELTA database to PAUP format and into distance matrices, for cladistic and phenetic analyses. Comments on the taxa included are provided, and an annotated and illustrated character list is presented. Subsets of characters and taxa were analysed and the results compared with previous classifications of the family, including one recently derived from manual cladistic analyses. The cladistic analyses were more successful in providing reasonable hypothetical phylogenies than in providing classifications permitting useful generalisation: The phenetically derived trees are in general similar to the cladograms, but they are more highly structured and correspond more closely to a previously published, manually derived cladogram. A suprageneric classification of the Cyperaceae is proposed, in which the genera are explicitly assigned to twelve tribes and two subfamilies. The interactive program INTKEY was used for preparing group descriptions and diagnoses, thus greatly facilitating comparisons among alternative classificatory solutions. Weaknesses of literature generalisations, particularly at higher taxonomic levels, highlight the need for more comparative data.
Carex vulpina and C. otrubae are difficult species to distinguish using floral characters. As an alternative, SEM of the nutlet surface, stomatal morphology and isozymes have been used as taxonomic tools to distinguish between the two taxa. The three methods show differentiation along species lines and are suitable to identify hybridisation. Stomatal morph-ology and isozymes reveal evidence of possible introgression between the two species.
Achenes of nine Indian Fimbristylis from sections Cymosae, Fuscae and Trichelostylis were treated with an acid solution to remove their cuticle and outer periclinal epidermal walls. Subsequent scanning electron micrographs revealed micromorphological characters of potential systematic merit. However, an attempt to correlate the achene micromorphology with the sectional ranks proposed by Kern, failed, with respect to the Cymosae and Trichelostylis. Such results suggest that either : (1) the sectional parameters recognized by Kern may be in need of revision or, (2) that the achene micromorphology is of little value in delimiting Fimbristylis sections.
Despite recent advances in molecular phylogenetic studies, deep evolutionary relationships in Cyperaceae are still not entirely resolved. Reduction of floral morphology and complex inflorescences pose difficulties to unravel relationships based on morphology alone. One of the most phylogenetically informative structures in Cyperaceae are the embryos. The utility of embryo characters and types in Cyperaceae systematics is reviewed in a molecular phylogenetic context using a DNA supermatrix incorporating sequences from five plastid (matK, ndhF, rbcL, rps16, trnL-F) and two nuclear ribosomal (ETS, ITS) regions. The phylogenetic hypothesis presented includes the most extensive sampling of the family to date. Fourteen qualitative morphological embryo characters were coded, ancestral state reconstructions were performed, and the embryo of each sampled genus was classified in a typological system based on key morphological features. Embryo morphology provides a valuable source of independent data for Cyperaceae systematics that can be used to place species with unknown affinities, when molecular data is not available, or when results of analyses are inconclusive or conflicting. Integrating embryo data will remain critical for future higher level studies of Cyperaceae evolution and classification.
Carex paracheniana, a new species of Cyperaceae from China, is described and illustrated. The new species is recognized as a member of Carex sect. Rhomboidales in having broadly ovoid perigynia and rhombic-ovoid nutlets. It is similar to C. cheniana and C. brevicuspis, but differs from the former in having narrower leaves 3–5 mm wide, staminate scales that are obtuse at the apex, and nutlets abruptly contracted into a ca. 1 mm long, slightly curved beak at the apex. It differs from C. brevicuspis in having ovate-elliptic pistillate scales that are glaucous or pale yellow and long-awned at the apex, and styles that are thickened at the base. Micromorphological characters of the leaf epidermis, perigynia, and nutlets are compared among taxa.
We described the achenes of 21 species of the genus Scleria reported in Costa Rica using 16 morphological characters and developed a key based only on achene characteristics. Specimens deposited in herbaria in Costa Rica were analyzed. We observed the achenes using a stereoscope and light microscope and took digital images that were used to measure the achenes. Besides, the achenes were observed using a Scanning Electron Microscope. A cluster analysis using achene characteristics was performed in order to know which species are morphologically similar. The intra-specific variation of the characteristics analyzed in the achenes studied is very small for all the species. Using characteristics of the achene, we could differentiate species among four of the five traditional sections of the genus used to classify the species: Hypoporum, Ophryoscleria, Schizolepis and Scleria. The key allows differentiating among 21 species of the genus Scleria previously reported in Costa Rica using only achenes. Besides the key, we prepared an illustrative guide for the genus using pictures taken with SEM and a stereoscope. The descriptions offer better information about the species that grow in Costa Rica.
All European taxa of Carex section Racemosae were assessed using macromorphological (mainly quantitative) and micromorphological characters (sculpture of utricles and nutlets). Morphological separation of the specimens examined was tested using discriminant function analysis (DFA). Micromorphology of utricles and nutlets was investigated using scanning electron microscopy (SEM). The results show that: (1) C. bicolor differs from other taxa in section Racemosae in macro- and micromorphological characters, justifying removal of the species from the section; (2) the morphological data support placing C. stylosa in a separate section, Stylosae; (3) morphologically, C. holostoma has a close affinity with taxa in subsection Papilliferae; (4) C. parviflora differs significantly from the remaining members of subsection Alpinae in the size and colour of the utricles and we suggest reconsidering the position of this species in subsection Alpinae; (5) morphological characters are insufficient for maintaining C. norvegica subsp. pusteriana and C. norvegica subsp. norvegica at subspecies level; (6) C. atrata, C. aterrima and C. caucasica are morphologically well-defined and valid species; (7) C. aterrima subsp. aterrima and C. aterrima subsp. medwedewii are morphologically similar and their taxonomic status requires further study, also at the molecular level.
Fuirena tenuis P.L. Forbes, a new species which has been found at a high altitude in the eastern Cape Province and Lesotho, is described and illustrated.
Aim
Across angiosperm families, the area occupied by a family is strongly correlated with its richness. We explore the causes of this area‐richness correlation using the cosmopolitan family, Cyperaceae Juss., as a model. We test the hypothesis that, despite a proposed tropical origin, temperate lineages in the family diversified at elevated rates. We test the hypothesis that the area‐richness correlation is maintained within intrafamilial clades, and that this relationship could be described as a function of niche space. We also test the hypothesis that the partitioning of geographical and ecological space, not the extent of this space, is the factor most closely associated with clade richness.
Location
Cosmopolitan.
Methods
We use molecular data from four genes sequenced in 384 taxa to develop a chronogram of Cyperaceae. We then develop a model of ancestral ranges and measure rates of diversification throughout the history of the family. Integrating data from over 4,800,000 digitized herbarium records, we characterize the range and niche of more than 4500 species and test for correlations of the species richness maintained within clades with range size, range partitioning, range overlap, niche, clade age and rate of diversification.
Results
Cyperaceae originated in South America in the late Cretaceous and subsequently dispersed throughout the globe. Of three increases in diversification rate, two occurred in the temperate Northern Hemisphere. The variable most closely associated with clade richness is the partitioning of geographical space by species within each clade.
Main conclusions
We show that species‐rich clades in Cyperaceae are not only more widespread, occupy more niche space, and diversify more quickly, but also exhibit patterns that are consistent with the partitioning of geographical and ecological space as a major correlate to diversification.
A data matrix was compiled to accommodate 38 macro- and micro-morphological characters of the achenes recorded comparatively for 12 Cyperus species (with three subspecies) using Light (LM) and Scanning Electron Microscopy (SEM).The data matrix was analyzed numerically using Jaccard’s coefficient as a similarity measure and the UPGMA clustering method. The recorded characters proved useful in two ways: (i) to delimit the taxa, and (ii) the dendrogram reflected the taxonomic relationships among the 15 taxa. Comparison between the taxonomic arrangement of the 15 taxa and previous classifications is provided.
Schoenoplectiella Lye, a genus segregated from Schoenoplectus (Rchb.) Palla, is accepted with expanded circumscription to include all the species formerly placed in Schoenoplectus sectt. Actaeogeton (Rchb.) J. Rayn. and Supini (Cherm.) J. Rayn. A revised description of Schoenoplectiella is provided and all the 50 species included in the genus are listed with 34 new combinations for the species, varieties, and hybrids. Two sections, Schoenoplectiella sectt. Schoenoplectiella and Actaeogeton (Rchb.) Hayasaka are recognized with 25 species for each, and a key to and descriptions for the sections are given. Morphological characters that define Schoenoplectiella and its sections, as well as those that distinguish the genus from allied genera, are discussed.
The cosmopolitan genus Carex L. represented about 1,800-2,000 species mainly in temperate and cold regions
of the world. 49 species of Carex found in North-eastern India and 33 species in the rest part of India.
Systematics of different taxa (subgenera, sections species pairs etc.) of the genus is quite complicated. In Carex,
utricle morphology and fruit epidermal silicabodies have been used to delimit species and sections. The present
study was undertaken to evaluate the taxonomic utility of these characters within sixteen species of Carex from
three subgenera and thirteen sections. The following sixteen species ( viz. Carex baccans Nees, C. myosurus
Nees, C. myosurus subsp. spiculata Boott, C. composita Boott, C. cruciata Wahlenberg, C. setigera D. Don, C.
breviculmis R. Brown, C. speciosa Kunth., C. insignis Boott, C. polycephala Boott, C. fusiformis Nees subsp.
finitima (Boott) Noltie, C. alopecuroides D. Don ex Tilloch and Taylor, C. teres Boott, C. longipes D. Don ex
Tilloch and Taylor, C. nubigena D. Don ex Tilloch and Taylor and C. remota L. subsp. rochebrunii (Franchet
and Savatier) Kükenthal) are evaluated in the present study. Perigynia of all the studied species are more or less
similar in their surface morphology. Surface appears smooth when examined with a dissecting microscope.
However, the epidermal cells have a distinctive morphology when viewed under higher magnification; differ
significantly in micromorphology (SEM-analysis). Perigynia have epidermal cells with thin convex/concave
outer wall that are collapsed in dried specimen. Variation in size, shape, color and number of nerve/costa and
texure of utricle are noticeable. Costal cells are narrower and more elongated, but also have thin, collapsed cell
wall. Several species have conspicuously nerved, stipitate perigynia others were not. Epidermal cells of achene
were polygonal to rhomboidal with little variation in size and shape. Epidermal cells of fifteen out of sixteen
species (except Carex nubigena D. Don ex Tilloch and Taylor) each have a single, rounded or nodulose silica
body with a basal platform. Although many unique characters were lacking, but interspecific variation in silica
bodies was consistent with the high level of morphological variation of utricle and previously detected
reproductive variation. The inclusion of micromorphological characters substantially enhances the resolving
power of macromorphologic characters in taxon analysis. These data indicated that utricle morphology and
micromorphological achene epidermal features (SEM features) are useful in identification of thirteen sections
and sixteen species of Carex.
Carex damiaoshanensis X. F. Jin & C. Z. Zheng, a new species of Cyperaceae from Guangxi, China, is described and illustrated. This new species resembles C. scaposa C. B. Clarke in its arundinaceous leaves and spikes arranged in a compound panicle, but differs from the latter in having glabrous culms, cauline leaves, and rachises of secondary inflorescences, panicle with fewer secondary inflorescences, and spikes with sparse pistillate flowers. Micromorphology of culms, cauline leaves, peduncles of secondary inflorescences, and achenes under SEM supports the recognition of this new species as well.
Small to tall perennial or annual herbs, rarely dwarf shrubs, shrubs, or lianas, terrestrial or helophytic, rarely aquatic, epiphytic, or epilithic; the perennial species tufted, rhizomatous, stoloniferous, bulbiferous, more rarely with a caudex. Cu/ms central, more rarely lateral, scapose or 1few-noded, more rarely many-noded, base sometimes bulbously thickened, (sub)trigonous to (sub)terete in cross-section, more rarely compressed, winged or 4–5–6-angular, rarely hollow or septate, smooth or scabrid, rarely viscid, papillose, or hairy. Leaves all (sub)basal or 1-few cauline, rarely all cauline, (spiro)tristichous, more rarely (spiro)distichous, lower leaves often (rarely all leaves) reduced to a sheath; sheaths usually closed, very rarely open, sometimes succulent or spongy; ligule present or absent, contraligule rarely developed; blade linear, rarely oblong, ensiform or terete, rarely pseudopetiolate. Inflorescence terminal, more rarely pseudolateral, paniculate, corymbose, spicate, anthelate, or capitate (often in combinations), more rarely reduced to a single spikelet, open or (partly) condensed, with few to many spikelets, branching from the axil of normal bracts, rarely from the axil of prophylls; prophylls present or absent; bracts leaflike or not, sheathing or not, lower primary bract sometimes culmlike and erect, spikelet bracts often small and glumelike. Spikelets with 1-many glumes, in cross-section rounded, 4–5-angular, laterally or rarely dorsiventrally compressed. Rachilla persistent or deciduous, rarely disarticulating into 1-flowered parts, rachilla internodes short or elongated, sometimes winged, or thickened and curved round the ripening fruit(s). Lateral spikelets usually prophyllate (with a basal, 2-keeled, usually empty glume), this prophyll rarely fertile, rarely utriculiform, and then enclosing the rachilla and its own flower. Glumes spirally arranged or distichous, rarely pentastichous, all glumes (sub)similar or of increasing length towards the (often upper) female or bisexual flowers, deciduous or persistent; all glumes fertile (often except for the lowermost and uppermost glumes), or only 1 to few upper (larger) glumes fertile, and then flower often enclosed by the wings of the next glume. Flowers bisexual or (functionally) unisexual, and then spikelets bisexual or unisexual, rarely dioecious. Hypogynous scales 0-(1-)3–6(-many), often bristle-like and retrorsely scabrid, more rarely laminar, sometimes with highly elaborated forms, shorter or longer than the fruit, usually deciduous with the fruit, rarely persistent on the rachilla. Stamens usually 3 (1 medio-anterior, 2 lateral), but not rarely reduced to 1 or 2, or 4–6 and even more numerouso; filaments sometimes strongly elongating after anthesis, rarely connate; anther basifixed, introrse, elliptic or oblong to linear, base often ± sagittate, connective often developed into a ± pronounced crest. Pistil superior, usually 3-carpellate, unilocular, ± trigonous, with 1 medio-anterior rib and 2 lateral ribs; style long, more rarely short, with 3 stigmatic branches, these sometimes with particular papillae; pistil sometimes 2-carpellate, and then either dorsiventrally or more rarely laterally compressed, rarely with a more elevated number of stigmatic branches. Style base distinct from the ovary or not, often conspicuously thickened and persistent, glabrous, scabrid, or variously hairy. Ovule single, basal, anatropous. Fruit an achene s.l. (nutlet), rarely drupaceous, usually ovate, elliptic, or obovate in outline, and (sub)trigonous to (dorsiventrally or, more rarely, laterally) lenticular in cross-section, surface smooth, punctate, reticulate or rugose, sometimes extremely differentiated, glabrous or finely hairy, sometimes seated on different types of cuplike structures. Seed with thin testa, free from the fruit wall, raphe and chalaza usually conspicuous, embryo small, basal, surrounded by abundant mealy or oily endosperm.
González‐Elizondo, M. S. & Peterson, P. M.: A classification of and key to the supraspecific taxa in Eleocharis (Cyperaceae) . – Taxon 46: 433–449. 1997. – ISSN 0040‐0262.
A comparison of former supraspecific classifications is made and a new, revised system is proposed for Eleocharis (Cyperaceae) . Four subgenera, seven sections, eight series, and seven subseries are recognized. A key to subgenera and sections is included along with type citations, synonymies, and representative species. Two new series, E. ser. Albidae and ser. Rostellatae , are described, along with one new combination, E. sect. Baeothryon , and two transfers in rank, E. subser. Acutae and E. subser. Sulcatae.
In Carex, vegetative anatomy and fruit epidermal silica bodies have been used to delimit species and sections, although frequent conflicts with traditional classifications have led many authors to question whether they can be used to infer evolutionary relationships. This conclusion, however, has largely been drawn from poorly circumscribed groups that have not been phylogenetically analyzed. This study was undertaken to assess the taxonomic and phylogenetic utility of these characters within Carex section Phyllostachys, a small (eight species), monophyletic group whose taxonomy and phylogeny has been extensively studied. Leaf and culm anatomy clearly separate the close species pair of Carex backii Boott and Carex saximontana Mackenzie, and they provide unique characters that distinguish Carex latebracteata Waterfall and Carex juniperorum Catling, Reznicek, & Crins. Anatomical and silica body characters strongly support the recognition of three species within the Carex willdenowii Willdenow complex (Carex willdenowii s.s., Carex basiantha Steudel, Carex superata Naczi, Reznicek, & B.A. Ford). Although unique characters were lacking in Carex jamesii Schw., infraspecific variation in its silica bodies was consistent with the high levels of genetic and morphological variation previously detected. Silica body characters support groups that are congruent with previous phylogenetic hypotheses derived from both morphological and molecular data. In contrast, vegetative anatomy, in this and previous studies, supports contradictory groups suggesting that its use in future phylogenetic studies below the sectional level in Carex may be limited.
The sectional delimitation of Carex section Rhomboidales sensu Kükenthal (1909) is problematic and, here, we provide contributions to a systematic understanding of nutlet micromorphology in section Rhomboidales. Nutlet micromorphology of 87 samples representing 71 taxa was investigated using scanning electron microscopy and nine nutlet types were recognized. Although some closely related taxa show similarities, nutlet morphology in section Rhomboidales is stable at the species level and is reliable for species identification. Based on the nutlet characters, section Rhomboidales is delimited and emended as having nutlet rhombic–ovoid, trigonous; beaks conspicuous, erect, rarely curved or coiled; periclinal walls of epidermal cells straight, with zero to two silica bodies. The species with nutlets of the C. harlandii-type and C. thibetica-type are included, and species of the C. chinensis-type and C. macrandrolepsis-type are ascribed to section Mitratae and section Infossae, respectively. Carex longirostrata and C. pseudolongirostrata (= C. nodaeana), formerly ascribed to section Careyanae or section Depauperatae, are well supported as members of section Rhomboidales. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 00, 000–000.
Eleocharis obtusa and Eleocharis ovata are recognized as distinct species or combined in recent taxonomic literature. To assess their morphological relationship and to evaluate all morphological characters potentially useful for their separation, 11 floral and achene characters were measured on 130 eastern Canadian herbarium specimens. Tubercle width was bimodally distributed and completely separated groups established on the basis of stamen number: plants referable to E. ovata had tubercles less than 0.5 mm wide and two stamens, whereas plants referable to E. obtusa had tubercles greater than 0.5 mm wide and three stamens. Since determination of stamen number requires dissection under a microscope and tubercle width requires accurate measurement, the most readily utilized character was found to be the ratio of tubercle width to achene width, which can be measured or estimated. The tubercle was less than 2/3 the width of the achene in E. ovata and more than 2/3 in E. obtusa. Differences in these characters were associated with significant but less dramatic differences in other characters, suggesting that the two taxa should be treated as distinct species. Keywords: Eleocharis obtusa, Eleocharis ovata, Cyperaceae, taxonomy, classification, Canada.
Eleocharis R. Br. achenes were examined employing scanning electron microscopy to ascertain the systematic potential of the achene wall. It was found that the epidermis has useful microscopic characters to assist in the systematic assessment of Eleocharis. Acid-treated achenes, with their cuticle and outer periclinal cell walls removed, revealed micromorphological differences in epidermal features among the 26 taxa studied. Characters of taxonomic interest include the configuration of the anticlinal cell walls, the contour of the cell lumens, in addition to the presence or absence of lumen pits, lumen depressions, and silica bodies. Such characters may be of value in assessing the infrageneric ranks of Svenson. Preliminary results support the Aciculares, Chaetarieae, Leiocarpeae, Multicaules, Ovatae, Palustres, Sulcatae, and Truncatae as natural taxa. In contrast, current data suggest that the Pauciflorae, Ocreatae, Rigidae, and Tenuissimae are unnatural entities. Key words: Cyperaceae, Eleocharis, SEM, achene, micromorphology.
The structures of the achene epidermal cells and their included silica deposits are illustrated and described for 28 species that have been referred to Carex section Hymenochlaenae. On the basis of these epidermal features, there are anomalous species in nearly all of the subsections and sections that have been circumscribed by Kükenthal, Mackenzie, Ohwi, and Koyama. Furthermore, some species with nearly indistinguishable achene surface features have quite different overall morphologies and have traditionally been placed in different sections. Similarities in the structure of the silica deposits in the achene epidermal cells may result from the retention of the primitive character state in several different lineages or from parallel development of modified silica bodies or wall structures in different lineages. Levels of homoplasy appear to be too high to make these characters reliable indicators of evolutionary relationships in Carex section Hymenochlaenae.
Carex yandangshanica C. Z. Zheng & X. F. Jin, a new species of Carex sect. Rhomboidales Kükenth. from Zhejiang, China, is described and illustrated. The new species is similar to C. thibetica Franch. and C. brevicuspis C. B. Clarke, but differs from the former by having trabeculate leaves, pistillate spikes with 6–10 staminate flowers or without staminate flower at the apex, and achenes at apex contracted into a slightly curved beak, and from the latter by having lateral culms, trabeculate leaves, and elliptic, rubiginous and awned pistillate scales. SEM microphotographs of perigynia and achenes are provided for the new species and related taxa.
Carex zhejiangensis X. F. Jin, Y. J. Zhao, C. Z. Zheng & H. W. Zhang, a new species of Carex sect. Rhomboidales sensu Kükenthal (Cyperaceae) from Zhejiang, eastern China, is described with illustrations. This new species is morphologically similar to C. filipes and C. macroglossa. Compared to C. filipes, C. zhejiangensis has elongated rhizomes, scattered culms, rather densely flowered pistillate spikes, achenes shortly stipitate at the base, and styles erect at the base. In contrast, C. filipes has short rhizomes, caespitose culms, sparsely flowered pistillate spikes, achenes obscurely stipitate at the base, and styles curved at the base. C. zhejiangensis differs from C. macroglossa by having elongated rhizomes and scattered culms, pistillate scales awned at apex, and perigynia contracted to a narrowly infundibuliform beak at apex. Diagnostic morphological characters, which are based upon population sampling, were analyzed in Carex zhejiangensis sp. nov., C. filipes and C. macroglossa.